Review of New Caledonian species of Oxyethira Eaton, with description of 17 new species, and new records for Hydroptila Dalman and Hellyethira Neboiss (Trichoptera, Hydroptilidae)

Abstract New Caledonian representation of the cosmopolitan genus Oxyethira Eaton is reviewed, with the description of new species bringing to 26 the total for the genus on the island. The species are referred to three subgenera: Trichoglene Neboiss (11 species), Pacificotrichia Kelley (13 species) and Dampfitrichia Ulmer (one species) and one species is unplaced to subgenus. A key is provided to Oxyethira species of New Caledonia. In addition, new records are given for two otherwise Australian species, Hydroptila losida Mosely and Hellyethira malleoforma Wells. Points marked on a series of small maps of New Caledonia indicate the site or sites at which the species were collected. This final paper in a series of generic revisions brings the hydroptilid fauna of the island of New Caledonia to 60 species, distributed in six genera.


Introduction
The cosmopolitan genus Oxyethira Eaton exhibits a diverse array of male genital structures and arrangements. The species show some variability in female terminalia, but exhibit extreme conservatism in larval and case morphology. Representatives of the genus found in New Caledonia appear to be no exception. This is apparent upon consideration of the variability among 26 species recorded here, and the similarity of the considerable number of unassociated final instar larvae collected at many sites by Mary (2002) in her survey of macroinvertebrates of the island's streams.
This final paper in a series of genus-level reviews Johanson 2012, 2014;Wells, Johanson and Mary-Sasal 2013) brings to 60 species, in six genera, the presently known hydroptilid fauna of New Caledonia, only surpassed by Ecnomidae in number of species found on the island (Espeland and Johanson 2010). The study is based for the greater part on collections of Hydroptilidae made by members of the Swedish Museum of Natural History and deals primarily with the genus Oxyethira. It also presents new records for the previously reported (Wells 1995) sole New Caledonian representatives of two other hydroptilid genera, Hydroptila losida Mosely, 1953 and Hellyethira malleoforma , both common in eastern Australia.
Oxyethira is well represented in New Caledonia, at genus level only surpassed in known species diversity among the island Trichoptera by the helicopsychid genus Helicopsyche (Johanson 1999;Johanson and Mary 2000), with 30 described species, and the ecnomid genus Agmina (Espeland & Johanson, 2010), with 28 described and nearly 50 undescribed species. Seven New Caledonian Oxyethira species were described by Kelley (1989), based on a collection in the B.P. Bishop Museum, Honolulu. He assigned the species to two subgenera: subgenus Trichoglene Neboiss (O. caledoniensis Kelley and O. insularis Kelley); and a new subgenus, Pacificotrichia Kelley (O. oropedion Kelley, O. dorsennus Kelley, O. indorsennus Kelley, O. melasma Kelley, and O. scutica Kelley), all assigned to the "oropedion group"). Oláh and Johanson (2010a) described three additional New Caledonia species: O. tompa, which they referred to subgenus Pacificotrichia; and two species, O. arok and O.
derek, which they assigned to subgenus Trichoglene. One representative of a third subgenus, Dampfitrichia Ulmer, the widespread SE Asian-Australasian species Oxyethira (Dampfitrichia) incana (Ulmer) described from Java, is recorded from New Caledonia for the first time.
Most of the 17 species newly described here from New Caledonian can be referred to the above three subgenera with a degree of confidence. One species, however, cannot be placed at present: O. macropennis sp. n. shares the diagnostic features of Oxyethira as defined by Kelley (1984) and is left unplaced.
Among females in the samples at least two general morphological forms can be recognised in abdominal terminalia: a short oviscapt of the form illustrated by Kelley (1989: figs 55, 56) for the Vanuatuan O. efatensis Kelley; and a slender, elongate oviscapt such as he illustrated for O. oropedion (Figs 50,51) and O. scutica (Figs 52,53), sometimes with a small, rounded black area ventrally on abdominal segment X as in O. oropedion. The distinctive female of O. incana is readily recognised by the quadrate black patch ventrally on segment X. Some of the other females were associated with males tentatively, but we are not sufficiently confident of their identity to include them here.
Apparent distributions of species are difficult to interpret (see . Almost all collections were made during November to mid January, normally the warmer season of the year. At least over that period, some species appear to be very localised, others widespread and still others disjunct in distribution, being taken from far northern and far southern localities. Further studies are needed at other times of year to determine whether these data reflect reality, seasonality, or some aspect of behaviour, such that collecting methods missed particular species.

Methods
Most of the material this study is based upon was collected in light traps and Malaise traps situated near or across running water. Specimens were prepared for study as Canada balsam slide mounts following maceration in KOH and clearing in clove oil. Male genitalia are illustrated in line drawings, traced from draft figures using Adobe Illustrator CS5, for species for which suitable slides are available.
An identification key and descriptions of New Caledonian Oxyethira species are provided, as well as brief diagnoses of previously described Oxyethira species and new illustrations of their male genitalia, drawn from types and/or newly collected non-type specimens. Species descriptions are based on male genital features, although identification of homologies among these is often difficult, especially for some of the more aberrant species. Usually diagnostic features are indicated on figures. Terms applied to genital structures follow the recommendations of Oláh and Johanson (2010b) and itemised by Wells and Johanson (2014: 3) when reviewing New Caledonian species of the genus Acritoptila.
Development of a useful key for easy identification of species was difficult as observation of most readily diagnostic features requires preparation of slide mounts of specimens and examination under a compound microscope.
Collection sites for species were plotted on a series of maps . Specimens in this study are deposited in the following repositories: MNHP Muséum National d'Histoire Naturelle, Paris, France NHRS Swedish Museum of Natural History, Stockholm, Sweden ANIC Australian National Insect Collection, CSIRO, Canberra, Australia BPBM Bishop Museum, Hawaii, USA

Oxyethira Eaton Subgenus Trichoglene Neboiss
The chief diagnostic characteristics that Kelley (1989) notes for subgenus Trichoglene are: in males, "a complete non-excised [abdominal] segment VIII", identified as plesiomorphic, and "aedeagus with recurved sub-distal spinous process and subgenital processes widely separated and partly fused with each pleuron of segment IX", features identified as apomorphic. Additional features are included in Kelley's description of the subgenus, including a titillator on the "aedeagus" [= phallic apparatus]. The subgenital processes in most members of this subgenus are in the form of a pair of well-separated rods, spines or strap-like structures, connected basally with the gonopods (= inferior appendages of Kelley) and a pair of digitiform membranous lobes, each bearing an apical seta.
In discussions of subgenus Trichoglene (Kelley 1984(Kelley , 1989, some confusion is evident in understanding of the type species of Trichoglene. Neboiss (1977) established the genus Trichoglene for Trichoglene columba Neboiss, described from Tasmania. This species was recognised by Wells (1981) as a species of Oxyethira. Upon designation of Trichoglene as a sugenus of Oxyethira, Kelley (1984) incorrectly gave the New Zealand Oxyethira albiceps (McLachlan, 1862) (= Hydroptila albiceps McLachlan, 1862) as the type. He repeated and compounded the error (Kelley 1989) by stating that "the type species of Trichoglene was incorrectly identified as Oxyethira columba (Neboiss) in Kelley (1984) [which it is not]… [i]t should be O. albiceps (MacLachlan)'. Perhaps he meant to imply that Hydroptila albiceps and Trichoglene columba (= Oxyethira columba) are synonyms, but they are distinct. Trichoglene columba Neboiss is the type species of the subgenus. Subgenus Trichoglene is Australasian in distribution, occurring in Australia, including Tasmania and Norfolk Island, and in New Zealand, as well as New Caledonia. Among New Caledonian species, Kelley 'E, 117 m, 20.xi-12.xii.2003, Malaise trap, loc#035, leg. K.A. Johanson (NHRS).
Etymology. Name spinifera is descriptive of the sharp spines of the subgenital processes.
Remarks. This species appears to be quite widespread from the far south towards the north of the island (Fig. 86).
Oxyethira (Trichoglene) tiwaka sp. n. http://zoobank.org/B1B6D5FA-BB85-4891- 75,87 Diagnosis. Readily recognised by the short, blunt, darkly sclerotised peg-like gonopods but in other respects showing very close resemblance to O. (T.) spinifera which has gonopods reduced so severely that they can be recognised as only small convexities on the apico-ventral margin of segment IX. Also similar to O. (T.) perignonica sp. n. in having abdominal segment IX subquadrate, but that species has the gonopods situated laterally, and curved mesally, and the subgenital processes in the form of convergent, rather than parallel, spines.
Male, genitalia . Abdominal segment IX quadrate in ventral view, slightly concave apico-ventrally, dorsally with proximal margin excised, V-shaped. Gonopods in form of two widely spaced, blunt black pegs. Setose lobes of ventral process angled laterally, ventral processes elongate, widely separated, tapered to narrowly rounded apices. Phallic apparatus elongate, with slender titillator and apically narrow, sinuous spine.
Etymology. Named for the river beside which one of the specimens was collected. Remarks. Oxyethira tiwaka was collected quite commonly in the southern region, but at only two disjunct localities in the north (Fig. 87). A photograph of the type locality with the trap is rendered in Fig. 75.
Male, genitalia (Figs 7-9). Abdominal segment IX in ventral view subquadrate, distally truncate with a small tuft of short setae each side of midline. Gonopods in ventral view forming a short, stout, mesally directed spur at each apico-lateral angle; setose lobes and rods of subgenital processes widely separated at bases, rods obliquely arranged, apically convergent; in lateral view, rods sharply down-turned. Phallic apparatus elongate, almost length of segments VII-IX, with a fine titillator and, subapically, a slender spine which in some specimens lies parallel to the length of the phallic apparatus, in others is twisted about it orthogonally. Material examined. Holotype. Male (on slide), New Caledonia, Province Sud, stream draining to Marais de la Rivière Blanche, 5 km SW Pont Pérignon, 22°09.513'S, 166°39.942'E, 180 m, 6-16.xi.2003, Malaise trap, loc#011, leg. K.A. Johanson (MNHP).
Etymology. Named for the bridge on the river near where the holotype was collected.

Remarks.
Taken only at several sites in the south of the island (Fig. 88), this species appears to have a highly localised distribution. A photograph of the type locality with the trap is rendered in Fig. 76. Etymology. abbreviata, named for the very abbreviated male genital structures.

Oxyethira
Remarks. This species was collected from several quite central sites (Fig. 89) from small rocky streams. A photograph of the type locality with the trap is rendered in Fig. 77. Diagnosis. Males resemble superficially those of O. perignonica both having mesally directed, laterally situated gonopods, but in ventral view these are more slender than those of O. perignonica and the rods of the subgenital processes are aligned in parallel with the distal margins of sternite IX in contrast to O. perignonica in which the they form sharp spines angled obliquely.
Etymology. The name incurvata is descriptive of the orientation of the gonopods.

Remarks.
From the collecting records the distribution of this species appears to be highly disjunct (Fig. 90), with records from the northern and southern extremes of the island. A photograph of the type locality with the trap is rendered in Fig. 78. 75,91 Oxyethira caledoniensis Kelley, 1989: 196, figs  Remarks. Delineation of this species among the large collection before us proved difficult, with only a very few specimens conforming closely to the holotype. In describing the species, Kelley (1989) had access to only a single specimen, re-examination of which shows it to be as illustrated in Kelley's fig. 56 (redrawn here from the type in Fig.  14), except that the subgenital processes are gently curved mesally, not slightly sinuous as figured by Kelley. A considerable number of specimens have been examined that agree in general features, but have abdominal segment IX either shorter but much the same shape as in the holotype male, or more elongate and rounded anteriorly; in some the spine on the phallic apparatus is longer and more strongly recurved and arising closer to the apex than in the type. This latter form is illustrated in Fig. 17 and was initially thought to be a separate species. However, following examination of further material of forms intermediate between O. caledoniensis sensu Kelley and this particular form, it is included tentatively as a variant form of O. caledoniensis together with all newly available specimens with the apico-ventral margin of abdominal segment IX truncate, sometimes with some slight marginal sclerotisation, the gonopods reduced to short domes, and abdominal segment IX sub-triangular to conical. Future studies may show reveal that these represent more than one species. Oxyethira caledoniensis is recorded from sites along the length of the island, but most commonly in the far south (Fig. 91). A photograph of one of the northern collecting sites is shown in Fig. 75 Antennae: male with 20-21 flagellomeres; flagellomeres quadrate to slightly rectangular in profile; female with 17 flagellomeres, terminal 2 pale, rest dark. Anterior wing length: male 1.2-1.8 mm (n=10); female 1.4-1.6 mm (n =10). Tibial spurs 0,3,4. Abdominal sternite VII with small median spur offset from distal margin. Female with length of abdominal segment IX almost twice width, distal margin, with a sclerotised margin, mesally produced distally. Segment X stouter at base than apex, gradually tapered distally, truncate apically.

Oxyethira (Trichoglene) caledoniensis Kelley
Material Oxyethira arok has been collected from disjunct localities in the far south and central part of the island (Fig. 92).
Oxyethira (Trichoglene) amieu sp. n. http://zoobank.org/B4BFF0C3-89A6-4825-A833-EFDFB92E5BD8 Figs 23, 24, 93 Diagnosis. Males are similar to O. arok and O. houailou sp. n. in the shape of abdominal segment IX, which in ventral view is strongly tapered and more or less triangular proximally, but both O. amieu sp. n. and O. houailou sp. n. also taper distally, while O. arok is more or less parallel-sided in distal half; in O. amieu sp. n. and O. arok gonopods are so reduced they cannot be identified clearly whereas in ventral view they are subquadrate in O. houailou sp. n.
Male, genitalia (Figs 23,24). Abdominal segment IX in ventral view subtriangular in proximal half, with proximal margin broadly rounded, distally tapered to about half maximum width; in lateral view triangular; gonopods reduced completely, subgenital processes rod-like, tapered distally, setal lobes almost at right angles to length of body; phallic apparatus with slender titillator and narrow, elongate subapical spine.
Male, genitalia . Abdominal segment IX in ventral view subtriangular in proximal half, with proximal margin truncate to broadly rounded, distally tapered to about half maximum width at distal margin; in lateral view triangular. Subgenital processes in form of stout sclerotised rods, their apices blunt, and short setose lobes, in ventral view almost at right angles to length of body. Phallic apparatus elongate,  Etymology. Named for the settlement of Houaïlou, near the type locality.
Remarks. This species was collected from widespread localities along the length of the island (Fig. 94).

Oxyethira (Trichoglene) insularis Kelley Figs 28-30, 95
Oxyethira insularis Kelley, 1989: 196, figs 34, 43, 57. Revised diagnosis. This species shares with O. parinsularis sp. n. the feature of gonopods, in ventral view, in the form of a prominent, sclerotised, Y-shaped structure, arising at around the distal third of abdominal segment IX, but differs from O. parinsularis sp. n. by having the fused basal section over 2× length of the divergent distal arms, not forming a shallow sclerotised band, and the bifid distal arms about 1/3 length of basal stem, not greatly exceeding length of base as in O. parinsularis sp. n.
Male antennae with 18 flagellomeres; flagellomeres rectangular in profile, without sensilla placodea, with abundant sensilla auricillica. Anterior wing length 1. Remarks. Among the extensive collection of New Caledonian Oxyethira at hand, only three specimens of this species were identified, all from the south of the island (Fig. 95). Other specimens from one of the sites at which they were taken are distinct and are referred to O. parinsularis sp. n. Description. Male antennae with 18-19 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea, with numerous sensilla auricillica; anterior wing length 1.5-1.8 mm (n=7); tibial spurs 0,3,4; abdominal sternite VII with median spine.

Oxyethira
Male, genitalia (Figs 31-34). Abdominal segment IX rounded proximally, in ventral view apically almost truncate but shallowly excavated medially, in dorsal view deeply and roundly excavated, with a pair of short, curved sclerotised processes mesally, interpreted as homologues of ventral processes, with associated short setose lobes; gonopods forked, forming a pair of widely divergent slender, curved spines, basally fused in a narrow band; phallic apparatus with a strongly recurved narrow spine apically; titillator present.
Material Remarks. The close similarity between this species and O. insularis is worrying, especially since both were taken at one site, yet the differences are clear and consistent. The species was taken only in the southern province (Fig. 96). A photograph of the type locality with the trap is rendered in Fig. 79.

Subgenus Pacificotrichia Kelley, 1989
Subgenus Pacificotrichia, based on the type species, Oxyethira oropedion Kelley, was diagnosed by Kelley (1989) by the following features of males: "… shallowly excised venter VIII …, the deeply excised dorsum VIII …, fused R4 and R5 forewing veins, configuration of the subgenital processes [these are fused], and reduction or loss of the pre-apical spur on the meso-tibia". Kelley commented on the similarity between the genitalia of males of this subgenus and those of the minima group in subgenus Dampfitrichia, but noted that in Pacificotrichia the subgenital processes are "distally fused … do not bear distal setae … and retain the bilobed process". The structure formed by the fused subgenital processes is generally characteristic for species, forming, in ventral view, what appears to be a plate ventral to the phallic apparatus. In common with minima group species in Dampfitrichia, most species in the Pacificotrichia group have a slender mid-ventral apodeme usually almost as long as the venter of abdominal segment IX, and unlike members of subgenus Trichoglene, lack a titillator on the phallic apparatus.
Seven species were referred by Kelley (1989) to subgenus Pacificotrichia, among which five were referred to an oropedion group and two to an efatensis group; the latter group was recorded only from Fiji and Vanuatu (one species each). However he did not define the two groups, and retention of these groups seems unnecessary.

Oxyethira (Pacificotrichia) oropedion Kelley
Revised diagnosis. Males are readily recognised by the short, setose mid-apicoventral lobes situated between the gonopods [present in the holotype, although not illustrated clearly by Kelley (1989: fig. 60)]. In other respects they resemble O. quadrata sp. n., O. indorsennus, and O. rougensis sp. n. but differ in ventral view from O. quadrata sp. n. in the apically subtriangular to rounded shape of the subgenital process, rather than quadrate, and the simple, straight phallic apparatus lacking distal flanges; from O. indorsennus in the wider separation of the gonopods; and from O. rougensis sp. n. by the broad, shallow excision of abdominal segment VIII, rather than deep almost V-shaped excision. The female is distinctive in having abdominal segments IX and X slender with  Remarks. Oxyethira oropedion is one of the more commonly collected New Caledonian Oxyethira species, occurring throughout the island (Fig. 97); several of the collecting sites are shown in photographs of type localities of other species (Figs 75, 76, 79, 81). Kelley's (1989)  Diagnosis. This species is distinguished in the male from the similar species O. oropedion by the broadly truncate subgenital process rather than triangular to rounded, flanges on the phallic apparatus; and by the significantly longer antennae of 32 flagellomeres compared to 22-24 flagellomeres.
Male, genitalia (Fig. 38). Abdominal segment VIII rounded proximally, tapered slightly to distal margin, a pair of small setose lobes mesally at base of gonopods, gonopods in form of short discrete conical lobes, setose lobes of ventral processes short, subgenital process a subquadrate plate. Phallic apparatus distally with a lateral flange, but no free spine or titillator present. Etymology. The name quadrata is descriptive of the shape of the subgenital processes.

Remarks.
Only one specimen of this species was identified, from the far south of the island (Fig. 98). Were it not for the exceedingly long antennae, we would probably have referred it to O. oropedion, broadening the concept of O. oropedion. A photograph of the type locality with the trap is rendered in Fig. 79. Kelley Figs 39,40,80,99 Oxyethira dorsennus Kelley, 1989: 199, figs 35, 44, 58. Revised diagnosis. Males are distinguished from the closely similar O. indorsennus by having spur formula 0,2,4, apical margins of gonopods truncate and subgenital process triangular rather than rounded as in O. indorsennus. These features also distinguish O. dorsennus from O. oropedion, which has apically well-separated gonopods with a pair of small setose lobes midventrally. Kelley (1989)  Remarks. Only three further specimens of this species have been identified, all from the south of the island (Fig. 99). They conform in detail with the type. A photograph of the type locality with the trap is rendered in Fig. 80.
Remarks. Kelley (1989: 199, fig. 36) distinguished O. indorsennus from O. dorsennus on the basis of difference in spur formula (0,3,4 cf. 0,2,4) and "… shape of dorsum VIII". However, examination of the types shows O. indorsennus to differ also in shape of the apical margins of gonopods and subgenital process, these being rounded as illustrated for O. tompa which also shares features such as spur formula and number of antennal flagellomeres (=24). Thus we are synonymising O. tompa with O. indorsennus. The species was collected widely throughout the island (Fig. 100).
This species shows a general resemblance to O. smolpela Wells, from New Guinea, but that species has a distinctive titillator on the phallic apparatus which is lacking in O. indorsennus.  O. quadrata, O. dorsennus, and O. indorsennus, but unlike any of those species, males have venter of abdominal segment VIII deeply and narrowly excised apico-medially and subgenital process sculpted latero-distally.
Male, genitalia. Abdominal segment VIII conical, distally about half width of proximal quarter. Segment IX in ventral view almond shaped, in lateral view triangular proximally, mid-dorsally less than half length of venter. Gonopods fused basally, discrete and rounded distally, with slender mid-ventral apodeme reaching to proximal margin of segment IX; subgenital processes fused, forming a stout plate, rounded distally in ventral view, angular in lateral view, with subapical sculpturing towards apex on each side, and small notch mid apically; setose lobes about 2/3 length of plate. Phallic apparatus slender, straight, with short apical spine.

Remarks.
Oxyethira rougensis was taken only at the type locality, in the north-west of the island (Fig. 101). A photograph of the type locality with the trap immediately below the small waterfall is rendered in Fig. 81.
Male, genitalia. In ventral view with abdominal segment VIII ovoid, apical margin with wide V-shaped excavation medially; without midventral apodeme; in lateral view, with apical margins broadly rounded; abdominal segment IX almond shaped. Gonopods sharply triangular; subgenital processes fused, forming subtriangular plate that is sharply triangular in lateral view; setose lobes less than half length of plate. Phallic apparatus extremely long, apically bifid, split into two equal-length sections.
Etymology. Named for Mt Mouirange near which the type was collected.
Male, genitalia (Figs 50-52). Abdominal segment VIII tapered slightly towards apex, only slightly longer than wide, with small cleft apico-ventrally and short apico-lateral lobes; in ventral view abdominal segment IX broadest mid length, tapered sharply proximally and distally, with gonopods forming a pair of narrow curved spines closely adpressed to an elongate triangular ventral process that terminates with a pair of tiny setae; phallic apparatus elongate, slender, with a long fine sinuous subapical process.
Remarks. The apparent distribution of this very distinctive species is very odd, with one sample being taken in the extreme north of the island, the rest at sites in the south-west (Fig. 103). A photograph of the type locality with the trap is rendered in Fig. 75.
Oxyethira (Pacificotrichia) enigmatica sp. n. http://zoobank.org/BFA21817-6E77-4056- BD89-45C2483C51F3 Figs 53-54, 82, 104 Diagnosis. In having an elongate apical spine on the phallic apparatus, males of this species resemble those of O. scutica, but in O. enigmatica sp. n. the spine is shorter and strap-like, not thread-like as in O. scutica; abdominal segment VIII tapers and gradually increases in width distally, and distal margin of venter is more widely and shallowly excavated than in O. scutica in which it is deeply and narrowly excised.
Male, genitalia (Figs 53-54). Abdominal segment IX in ventral view wider distally than proximally, with distal margin shallowly excavated, proximal margin rounded, midventral apodeme present; in lateral view dorsal margin about half depth of ventral margin. Gonopods fused basally, distally discrete, conical; subgenital processes forming short, subquadrate plate. Phallic apparatus straight, with long sharply twisted apical spine, at right angle to and almost one third length of phallus; without titillator.
Etymology. Enigmatica: enigmatic in presenting a puzzle, being so similar to O. (P.) scutica in some respects, yet distinctive.
Remarks. This species is known only from the large sample taken at the type locality in the extreme south of the island (Fig. 104). A photograph of the type locality with the trap is rendered in Fig. 82. 105 Oxyethira melasma Kelley, 1989: 200, figs 37, 45, 59. Note. A mismatch between the published description and Kelley's (1989) figures and the holotype specimen in the BPBM labelled "Oxyethira melasma" is a puzzle. The genital features of the holotype (examined by AW) are as in Fig. 56, and disagree with Kelley's text description and illustrations (his figs 37, 45, 59) of "Oxyethira melasma". However, the tibial spur count of the holotype is 0,2,4, as given by Kelley, and as in the type species. Kelley's three figures of genital structures agree with the features of a series of specimens, described here as Oxyethira nehoue and illustrated in Figs 58-60; in contrast to Kelley's "holotype", these specimens all have a tibial spur count of 0,3,4.

Oxyethira (Pacificotrichia) melasma Kelley
Oxyethira melasma Kelley is here redescribed and figured from the holotype specimen. We can only suggest that Kelley had several specimens at hand, macerated one and labelled it "holotype", but described and illustrated a specimen other than that labelled "Type". The type has to be the name-bearing specimen and is redescribed here.
Revised diagnosis. Males resemble O. nehoue sp. n., O. spicula sp. n., and O. ouenghi in having have a more or less triangular median ventral processes in the male genitalia. However, they most closely resemble O. nehoue sp. n. from which they are distinguished by spur count 0,2,4, gonopods fused, in ventral view sharply tapered proximally, narrowly parallel-sided distally, rounded apically, apex of phallic apparatus rounded without apical spine, in contrast to O. nehoue sp. n. in which the fused gonopods appear triangular in ventral view, and acuminate apically, and ventral process in lateral view sclerotised and arched ventrally.
Male, genitalia (Figs 55-57). Abdominal segment VIII more or less conical, a deep broadly V-shaped excision apico-ventrally. Abdominal segment IX in lateral view broadly bell-shaped; in ventral view rounded proximally, tapered distally, a pair of small short processes apically, each bearing a single seta. Gonopods sclerotised, fused, tapered to narrowly truncate apex, basal setose processes widely separated, slender, elongate, but shorter than fused gonopods, a slender basal apodeme midventrally; subgenital plate broad based, tapered to rounded apex, mostly membranous, but with a small ventrally curved prominence subapically. Phallic apparatus swollen in basal third, narrow in distal 2/3, without apical spine, ejaculatory tube medial.
Remarks. The species was taken at a number of sites in the southern province of the island (Fig. 105).
Oxyethira (Pacificotrichia) nehoue sp. n. http://zoobank.org/EE2F9749- F988-4653-9556-D40FFD285635 Figs 58-60, 83, 106 Diagnosis. This species resembles O. melasma and was illustrated and described as that species by Kelley (1989); however the holotype is as in Figs 55-57, see discussion above. The two species are distinguished by male genital features: O. nehoue sp. n. has abdominal segment VIII in ventral view shallowly and narrowly excavated mid apically, not widely and deeply; fused gonopods tapered to an acute apex, not truncate; subgenital process elongate rectangular, truncate apically, not rounded, in lateral view sclerotised and arched ventrally; and phallic apparatus with a broad spine arising sub apically.
Genitalia (Figs 58-60). Abdominal segment VIII in ventral view rounded proximally, gradually tapered towards apex, with a shallow, narrow excavation mid apically. Abdominal segment IX similar in shape to VIII. Gonopods fused, triangular in ventral view, dorsal setose lobes slightly shorter than gonopods conjoined at base, subgenital process elongate, forming narrowly rectangular plate, apically truncate, but with slight bulge mid-apically. Phallic apparatus with a broad spine arising subapically, extending beyond apex.
Female unknown.
Etymology. Named for the Rivière Néhoué where it was collected.
Remarks. The distribution of this species is similar to that of O. ouenghi, mainly collected from a cluster of southern sites, but with one site in the far north (Fig. 106). A photograph of the type locality with the trap is rendered in Fig. 83.

Oxyethira (Pacificotrichia) scutica Kelley Figs 61-63, 107
Oxyethira scutica Kelley, 1989: 200, figs 39, 47, 52, 53, 61. Diagnosis. Males superficially resemble those of O. enigmatica, having a long apical process on the phallic apparatus, antennae with 23-25 flagellomeres, and spur formula of 0,2,4, but are distinguished by apical process on phallic apparatus longer and thread-or whip-like in appearance, rather than strap-like as in O. enigmatica, and abdominal segment VIII with distal margin of venter of deeply and narrowly excised, compared with shallowly excavated margin of O. enigmatica. Females resemble those of O. oropedion (Kelley 1989: figs 50, 51, both having abdominal terminalia in form of a slender oviscapt and cerci slender, elongate, about length of segment X, but females of O. scutica lack the darkly sclerotised area on the venter of segment IX (Kelley 1989: figs 52, 53).
Remarks. This appears to be another southern species (Fig. 107).
Oxyethira ( which have more or less triangular median ventral processes in the male genitalia, but can be recognised by the expanded apex of the phallic apparatus with a prominent acute spine, the very long, proximally rounded, abdominal segment VIII that tapers distally and completely obscures segment IX, and the shape of the plate formed from fused gonopods and subgenital processes. Description. Male antennae damaged, at least with 19 flagellomeres, flagellomeres rectangular in profile, without sensilla placodea; anterior wing length 1.9 mm (n=1); tibial spurs 0,3,4; abdominal sternite VII with a coarse spur medially.
Male, genitalia (Figs 64-66). Abdominal segment VIII pear-shaped, ventrally with narrow distal margin excavated apically; abdominal segment IX obscured by VIII; gonopods fused forming a triangular plate ventrally with subgenital process a short knobshaped lobe apically and a deep mid ventral apodeme reaching almost full length of segment; phallic apparatus elongate and slender in proximal 3/4, irregularly dilated distally, with sharp apical spine and short lateral process.
Female unknown, although a single unknown female of an Oxyethira species was collected with the holotype and could be of this species. The terminalia of this female are slender and elongate, of the form seen in O. oropedion and O. scutica, with a Vshaped marking ventrally on abdominal segment VIII.
Etymology. Name being descriptive of the spicule-like spine on the phallic apparatus.
Remarks. This species is known only from the type specimen, collected in the far south (Fig. 108) and now on a prepared slide. It is impossible to know if the state of abdominal segment IX of the type is the usual situation, or simply unusual that segment has retracted in this particular specimen. Regardless, the form of the phallic apparatus is highly distinctive. Diagnosis. One of the New Caledonian species of Oxyethira with genitalia retracted within the very rounded abdominal segment VIII but differs from other species with this feature such as O. incana and O. spicula by well-defined clearly branched gonopods and the phallic apparatus a simple rod, sharply bifid apically.
Male, genitalia (Figs 67-69 Etymology. Named for the finger-like lobes of the gonopods in male.

Remarks.
Oxyethira digitata shares a southern distribution (Fig. 109) with O. melasma and O. scutica. A photograph of the type locality with the trap is rendered in Fig. 84.

Subgenus Dampfitrichia Ulmer
Erected at genus level by Mosely (1937: p.169), and synonymised with Oxyethira by Ross (1944), Dampfitrichia was accorded subgenus status by Kelley (1984) in Oxyethira  (Ulmer), ventral view and phallic apparatus 72-74 O. macropennis sp. n., ventral and lateral views and phallic apparatus. Abbreviations: gon = gonopod; sp = subgenital process; st lb = setose lobe of subgenital process; VIII, IX = abdominal segments VIII and IX. and diagnosed as "… characterised by a fusion of veins R4 and R5 in the forewing and the subdistal sclerotised bridge between the subgenital processes"; Kelley noted that the phallic apparatus usually lacks a titillator. Diagnosis. Males of this species are distinguished from others in the New Caledonian fauna by abdominal segment VIII with disto-lateral angles spiny, and venter produced and rounded distally not excised apico-ventrally, forming a shield over other genital structures which are strongly fused; and by phallic apparatus curiously stout and medially curved, lacking a titillator. Female terminalia longer and terminally more slender than those of caledoniensis group species, but not as slender as in members of subgenus Pacificotrichia, with sternite X bearing a jet black quadrate plate.

Remarks.
In New Caledonia this species was taken only in the northern province (Fig. 110); elsewhere it is widespread from Java, through South-East Asia to New Guinea and northern Australia.
One highly aberrant species, O. macropennis sp. n., is here unplaced to subgenus. Males share the diagnostic features of species of Oxyethira, including wing shape and venation, but have unusual male genital features, possibly aligning the species more with species of Paroxyethira. For the present we assign it to Oxyethira, albeit tentatively.
Oxyethira macropennis sp. n. http://zoobank.org/3FA585C6-F464-4B22- B39E-C3B1172F5A0C Figs 72-74, 85, 111 Diagnosis. Immediately recognised by the remarkable form of the phallic apparatus, which has a contorted twist towards the stout seta-bearing apex, and the brush of shorter stout setae medially on abdominal sternite IX. By these features it is distinguished clearly from all other New Caledonian species.
Male, genitalia (Figs 72-74). Abdominal segment VIII quadrate in profile; abdominal segment IX subrectangular, ventrally bearing a brush of stout setae medially, and on each side a cluster of shorter setae subapically, mid dorsally short, apical margin shallowly excavated; gonopods may be represented by the sclerotised apico-lateral lobes on abdominal segment IX; phallic apparatus strongly twisted in distal half, beyond a short lateral process that may represent titillator, subapically irregular in shape, bearing a stout seta.
Remarks. Oxyethira macropennis is quite unlike all other New Caledonian species, however it conforms with the diagnostic features of members of the Oxyethira and thus is assigned to this genus, albeit somewhat tentatively.
Etymology. Name descriptive of the relatively large phallic apparatus.